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作物学报 ›› 2020, Vol. 46 ›› Issue (11): 1678-1689.doi: 10.3724/SP.J.1006.2020.04036

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

白菜型油菜粒色主效基因BrTT1的调控机制分析

王艳花1,2(), 荐红举1, 邱晓2, 李加纳1,*()   

  1. 1 西南大学农学与生物科技学院 / 油菜工程研究中心, 重庆 400715
    2 萨斯喀彻温大学, 加拿大萨斯卡通 S7N5A8
  • 收稿日期:2020-02-17 接受日期:2020-07-02 出版日期:2020-11-12 网络出版日期:2020-07-15
  • 通讯作者: 李加纳
  • 作者简介:E-mail:hawer313@163.com
  • 基金资助:
    本研究由中国博士后科学基金面上项目(2019M653319);重庆市自然科学基金博士后科学基金项目(cstc2019jcyj-bsh0102);高等学校学科创新引智基地项目(“111”项目)(B12006)

Regulatory mechanism of the seed coat color gene BrTT1 in Brassica rapa L.

WANG Yan-Hua1,2(), JIAN Hong-Jiu1, QIU Xiao2, LI Jia-Na1,*()   

  1. 1 College of Agronomy and Biotechnology, Southwest University / Chongqing Engineering Research Center for Rapeseed, Chongqing 400715, China
    2 University of Saskatchewan, Saskatoon S7N5A8, Canada
  • Received:2020-02-17 Accepted:2020-07-02 Published:2020-11-12 Published online:2020-07-15
  • Contact: Jia-Na LI
  • Supported by:
    This study was supported by the Project of China Postdoctoral Science Foundation(2019M653319);the Project of Chongqing Natural Science Foundation Postdoctoral Science Foundation(cstc2019jcyj-bsh0102);Project of Intellectual Base for Discipline Innovation in Colleges and Universities (“111” Project)(B12006)

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摘要：

白菜型油菜(Brassica rapa L., 2n = 20, AA)为十字花科芸薹属作物, 属于栽培油菜基本种。我国是白菜和白菜型油菜的起源中心, 与甘蓝型油菜相比, 其起源和栽培历史悠久, 遗传资源丰富, 具有天然而稳定的黄籽资源。大黄油菜是白菜型油菜中一种天然的黄籽资源, 其粒色鲜黄, 黄籽性状能够稳定遗传, 且具有大粒、含油量高、自交亲和性好等优点, 是研究油菜粒色性状的良好材料。本研究对白菜型油菜粒色主效基因BrTT1进行了进一步的功能验证, 并对BrTT1的粒色调控机制进行了初步解析。序列比较结果表明, BrTT1在不同遗传背景白菜型油菜黄籽、红褐籽、黑籽品种中存在固定的差异位点, 同一粒色材料所得BrTT1序列一致, 可用于预测粒色表型; 酵母双杂交分析表明, BrTT1可以与另外2个转录因子R2R3-MYB (BrTT2)和WD40 (BrTTG1)以及一个催化酶(BrTT3)相互作用。qRT-PCR结果表明, 超量和干扰BrTT1的表达, 导致类黄酮合成路径结构基因及其他关键调节基因BrTT2、BrTTG1的异常表达或不表达, 从而阻碍了原花色素的正常积累。研究结果进一步明确了BrTT1在白菜型油菜粒色形成中的调节活性。

关键词: 白菜型油菜, BrTT1, 类黄酮代谢, 酵母双杂

Abstract:

Brassica rapa (B. rapa L., 2n = 20, AA) is a specie of Brassica genus, belonging to the basic species of cultivated rapeseed. China is the original center of Chinese cabbage and Brassica campestris. Compared with Brassica napus, it has a long history of origin and cultivation and rich genetic resources, which has natural and stable yellow seed resources. Dahuang has the natural yellow seed resource in B. rapa. Its seed coat color is bright yellow, the yellow seed trait can be stably inherited, and Dahuang has the advantages of large grain, high oil content and good self-adhesiveness. Sequence comparison showed that nucleotide polymorphisms were solely found in BrTT1 sequences from different seed color lines (yellow, red or brown, and black), which could be used to predict seed color phenotype. Yeast two-hybrid analysis indicated BrTT1 could interact with two other transcriptional factors R2R3-MYB (BrTT2) and WD40 (BrTTG1), and one catalytic enzyme (BrTT3). Quantitative RT-PCR analysis of transgenic B. rapa lines with the gene down-regulated by RNA interference and up-regulated by overexpression revealed that two contrasting groups of genes were regulated by BrTT1 in the biosynthesis and deposition of flavonoids pigments in the seed of B. rapa. These results further define the regulatory activity of BrTT1 in seed coat color formation in Brassica species.

Key words: Brassica rapa, BrTT1, flavonoid, yeast two-hybrid

附表1

不同来源的白菜型油菜黄、红、黑籽材料"

编号
No.		 材料名称和来源
Material name and origin		 种子粒色表型
Phenotype of the seeds		 编号
No.		 材料名称和来源
Material name and origin		 种子粒色表型
Phenotype of the seeds
Y1 大黄(中国青海)
Dahuang (Qinghai, China)		 纯黄籽
Yellow seeds		 R6 18B167-168 (中国青海)
18B167-168 (Qinghai, China)		 红褐籽
Red or brown seeds
Y2 18B07-08 (中国青海)
18B07-08 (Qinghai, China)		 纯黄籽
Yellow seeds		 R7 18B171-172 (中国青海)
18B171-172 (Qinghai, China)		 红褐籽
Red or brown seeds
Y3 彭波黄(中国青海)
Pengbo yellow (Qinghai, China)		 纯黄籽
Yellow seeds		 R8 18B141-142 (中国青海)
18B141-142 (Qinghai, China)		 红褐籽
Red or brown seeds
Y4 18B001-002 (中国青海)
18B001-002 (Qinghai, China)		 纯黄籽
Yellow seeds		 R9 18B51-52 (中国青海)
18B51-52 (Qinghai, China)		 红褐籽
Red or brown seeds
Y5 Sarson (印度)
Sarson (India)		 纯黄籽
Yellow seeds		 B1 浩油 11号(中国青海)
Haoyou 11 (Qinghai, China)		 纯黑籽
Black seeds
Y6 18BCan01 (加拿大)
18BCan01 (Canada)		 纯黄籽
Yellow seeds		 B2 芦花小油菜(中国青海)
Luhua rapeseed (Qinghai, China)		 纯黑籽
Black seeds
R1 18B017-018 (中国青海)
18B017-018 (Qinghai, China)		 红褐籽
Red or brown seeds		 B3 街子小油菜(中国青海)
Jiezi rapeseed (Qinghai, China)		 纯黑籽
Black seeds
R2 18B021-022 (中国青海)
18B021-022 (Qinghai, China)		 红褐籽
Red or brown seeds		 B4 18B077-078 (中国青海)
18B077-078 (Qinghai, China)		 纯黑籽
Black seeds
R3 18B105-106 (中国甘肃)
18B105-106 (Gansu, China)		 红褐籽
Red or brown seeds		 B5 麻玉小油菜(中国青海)
Mayu rapeseed (Qinghai, China)		 纯黑籽
Black seeds
R4 18B131-132 (中国青海)
18B131-132 (Qinghai, China)		 红褐籽
Red or brown seeds		 B6 18B161-162 (中国青海)
18B161-162 (Qinghai, China)		 纯黑籽
Black seeds
R5 18B165-166 (中国青海)
18B165-166 (Qinghai, China)		 红褐籽
Red or brown seeds		 B7 18BCan02 (加拿大)
18BCan02 (Canada)		 纯黑籽
Black seeds

附表2

用于载体构建的引物序列"

引物名称
Primer name		 正向引物
Forward primer (5°-3°)		 反向引物
Reverse primer (5°-3°)
Yb-F/-R CCGGAATTCATGTTTTCATCACTCTCAAACCACT CGCGGATCCTTAAAAGTGCGTTTCAGAGACAGA
T2-F/-R CCGGAATTCATGATGAGAAAGAGAGAAAGTAGTA CGCGGATCCCTAACAATTAAAGTCCCAGAGACAA
T3-F/-R CCGGAATTCATGATTCTGGAGGAGAAAGATGCAT CCCATCGATTTAGCTATCTGAACGTTTTGGATCG
T7-F/-R CCGGAATTCATGGTGATGGGTACACCGTCTTCGT CGCGGATCCTCAAACAGGAACGCTGTGCAAGACC
T8-F/-R CCGGAATTCATGGATGAATTAAGTATTATACCGT CGCGGATCCCTAGAGTTTATTATTATATATGATT
T18-F/-R CCGGAATTCATGGGTAGCAAGATGCTGTTCAGTT CGCGGATCCTTAGCTTGCTTCTGGCAATGTGATC
TG1-F/-R CCGGAATTCATGGACAACTCAGCTCCGGACTCCT CGCGGATCCTCAAACTCTAAGGAGCTGCATTTTG
TG2-F/-R CCGGAATTCATGGAGGTGAAAGAGAGTAAGAGAG CGCGGATCCTCAAATGGCTTGATTAGAATGTTGT
T10-F/-R CCGGAATTCATGACAAGCACTGATCAGACCACCG CGCGGATCCTTAGGCCCATCGGTTTGTCTTAAAG
T12-F/-R CCGGAATTCATGGGGAGGAAGACGTGGTTCGACG CGCGGATCCTCAGAAGAGATAATCAAGTATCTGA
T5-F/-R CCGGAATTCATGAGACATTCTTGCTGTTATAAAC CGCGGATCCCTAAAGGGACTGACCAAAAGAGACA
F/R-TT1 ATGTTTTCATCACTCTCAAACCACT TTAAAAGTGCGTTTCAGAGACAGA
gCb-F/-R CACCATGTTTTCATCACTCTCAAACCACT AAAGTGCGTTTCAGAGACAGA
F35S3ND/CB-R GGAAGTTCATTTCATTTGGAGAG AAAGTGCGTTTCAGAGACAGA
FBTT1I GGATCCGACGTCGAGATCTATAAACATTCCC

附表3

用于差异表达分析的引物序列"

引物名称
Primer name		 正向引物
Forward primer (5°-3°)		 反向引物
Reverse primer (5°-3°)
28S ATAACCGCATCAGGTCTCCAAG CCTCAGAGCCAATCCTTTTCC
BrTT1 TCGCTACAACAATCTTCAGATGCACA TCCTGCACCCTTCAACGCAGC
BrTT2 AGCTGGTCTCAAGAGGTGTGGCA AGCCTCCCAGCTATCAACGACC
BrTT3 CAGGATGGATGTATTTCATGTCG TGTGCCTCGTTACGAGTGATAG
BrTT4 GACTACTACTTCCGCATCACCAACAG GCCTAGCTTAGGGACTTCAACAACC
BrTT5 CTTCCTCGGTGGCGCAGGTG ACACAGTTCTCCGTTACTTTCTCTGA
BrTT6 CGAGAAAGAGGCACTCACCAATG TGAACCTCCCGTTGCTCAGATA
BrTT7 GTGGTTGCCGCCTCTAAATC CTAGCTCGCGCATGAGTGTTC
BrTT8 GGCTGAAGAGGCTGCGTCGG GTGCTGTGCAAGCCCTCGCT
BrTT10 GCGACTGTGCCAAGAAACGGT CCCCACGTGAGATGTCTATCAAAGTG
BrTT12 GCTCCACAGAGACATACGAGCCG ACGGTGACGAAGCTGAGCATGTA
BrTT16 TGCTCACATCGGTCTCATCGTCT GCTCGTGTGGAGGAATGGAGGC
BrTT18 AAGAAGAGCTCGAGAGCATCAAC GCATACTTCTCCTTCTCTTCCAC
BrTTG1 TCCTCCGGCGACTTCCTCCG GCTGCGTCTCCACCACGGAC
BrTTG2 AAACCTAAAGCAAAGCTTGTCTCCCA ACTTCCTTTGACTTGCTTCTGTCCGT
BrBAN TTAACTGGGCATACCCAATCTC TGCATTTCTTTCCGGGTAATC

图1

BrTT1同源基因氨基酸序列比对"

图2

不同粒色的BrTT1基因的核苷酸序列比对 A: 不同粒色的白菜型油菜种子表型(黑色、红褐色和黄色); B: 来自不同粒色的 BrTT1的核苷酸序列。TT1-B: 从黑籽中扩增得到的BrTT1-B序列; TT1-R: 从红褐籽种子中扩增得到的BrTT1-R序列; TT1-Y: 从黄籽中扩增得到的BrTT1-Y序列。核苷酸多态性通过颜色突出显示, 共有序列下方的线代表内含子区域。标尺 = 5000 μm。"

图3

BrTT1基因在3种不同粒色种子形成期的表达分析 B1: 黑色种子系; R2: 红褐籽种子系; Y2: 黄籽种子系。DAF: 开花后天数。"

图4

白菜型油菜BrTT1的互作蛋白预测分析"

表1

预测白菜型油菜BrTT1的互作蛋白的信息"

基因ID
Gene ID		 蛋白类型
Protein type		 打分值
Score		 基因ID
Gene ID		 蛋白类型
Protein type		 打分值Score
Bra027457 BrTT3 0.996 Bra005210 BrTTG2-A, WRKY44 0.814
Bra035532 BrTT2 0.992 Bra023112 BrTTG2-B 0.814
Bra009770 BrTTG1 0.909 Bra026408 Uncharacterized protein 0.607
Bra037887 BrTT8 0.894 Bra013652 BrTT18 0.568
Bra009312 BrTT7 0.885 Bra016108 Uncharacterized protein 0.568
Bra040822 Uncharacterized protein 0.882 Bra009101 Uncharacterized protein 0.568
Bra003361 BrTT12 0.881 Bra039487 Uncharacterized protein 0.434
Bra007142 BrTT5 0.822 Bra035364 Uncharacterized protein 0.434
Bra037510 BrTT10-2 0.821 Bra006205 Uncharacterized protein 0.379
Bra020720 BrTT10-1A 0.821 Bra021170 Uncharacterized protein 0.365

图5

酵母双杂筛选BrTT1可能的互作蛋白 酵母细胞涂布于-Leu/-Trp/-Ade/-His的培养基, pGADT7- T/pGBKT7-53作为阳性对照, pGADT7-T/pGBKT7-laminC作为阴性对照。X-α-gal的染色深度表示互作强弱。"

附图1

PCR扩增检测RNAi BrTT1得到的阳性转基因植株 检测引物为F35S3ND/FBTT1I (附表2), 检测片段1.0 kb。+表示阳性对照。"

附图2*

PCR扩增检测过表达 BrTT1得到的阳性转基因植株 检测引物为F35S3ND/FBTT1I (附表2), 检测片段1.0 kb。+表示阳性对照。"

图6

过量和干扰表达BrTT1基因 A: 转基因种子中BrTT1的差异表达; O1~O4: 过量表达转基因株系; Ri1~Ri4: 干扰表达转基因株系; B: 非转基因黑籽材料; DAF: 开花后天数。B: 左、中、右分别为过量表达、干扰表达BrTT1基因、非转基因褐籽对照成熟种子表型。标尺 = 5000 μm。"

图7

RNAi和超量表达BrTT1基因的转基因株系中类黄酮相关基因的表达分析 O1~O4: 过量表达转基因株系; Ri1~Ri4: 干扰表达转基因株系; DAF: 开花后天数。"

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